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10 Bodnar Et Al Pg141a160 COR.Indd Rev. bras. paleontol. 18(1):141-160, Janeiro/Abril 2015 © 2015 by the Sociedade Brasileira de Paleontologia doi: 10.4072/rbp.2015.1.10 VOLTZIALES Y PINALES (= CONIFERALES) DE LA FORMACIÓN CORTADERITA (TRIÁSICO MEDIO), ARGENTINA, Y SU IMPLICANCIA EN LA RECONSTRUCCIÓN DE LAS CONÍFERAS TRIÁSICAS JOSEFINA BODNAR División Paleobotánica, Facultad de Ciencias Naturales y Museo, CONICET, Paseo del Bosque s/n, B1900FWA, La Plata, Buenos Aires, Argentina. [email protected] DANIELA PAULA RUIZ División Paleobotánica, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, CONICET, Av. Ángel Gallardo 470, C1405DJR, Ciudad Autónoma de Buenos Aires, Argentina. [email protected] ANALÍA EMILIA ARTABE División Paleobotánica, Facultad de Ciencias Naturales y Museo, CONICET, Paseo del Bosque s/n, B1900FWA, La Plata, Buenos Aires, Argentina. aartabe @fcnym.unlp.edu.ar EDUARDO MANUEL MOREL División Paleobotánica, Facultad de Ciencias Naturales y Museo, CIC, Paseo del Bosque s/n, B1900FWA, La Plata, Buenos Aires, Argentina. [email protected] DANIEL GANUZA División Paleobotánica, Facultad de Ciencias Naturales y Museo, Paseo del Bosque s/n, B1900FWA, La Plata, Buenos Aires, Argentina. [email protected] ABSTRACT – VOLTZIALES AND PINALES (= CONIFERALES) FROM CORTADERITA FORMATION (MIDDLE TRIASSIC), ARGENTINA, AND THEIR IMPLICATION IN THE RECONSTRUCTION OF TRIASSIC CONIFERS. In this work, fossil wood, leaves, and branches designated to the subclass Pinidae (Voltziales and Pinales) are described from Cortaderita Formation (Middle Triassic). Fossil materials were collected at Cortaderita and La Tinta creeks, at Barreal area from Barreal-Calingasta basin, Pre-Andean region of San Juan, Argentina. Leaves corresponding to three species of Heidiphyllum (order Voltziales), leafy twigs assigned to Elatocladus planus (Pinales) and permineralized trunks and branches ascribed to Cupressinoxylon zamunerae sp. nov. (Pinales), were recognized. The last two species exhibit characters that allow designating them to family Cupressaceae sensu lato. At the same time, it is demonstrated that they belong to the same plant, which is named as the new species of Cupressinoxylon here described. This reconstruction is based on kindred reinforcement and mutual occurrence criteria, besides morphological and anatomical characters. Thus, Cupressinoxylon zamunerae sp. nov. is interpreted as a tree 13 m tall, highly branched, with penultimate order shoots with scale-like leaves and last order shoots with alternate, sessile, single-veined, linear to acicular leaves distributed helically. These records confi rm the presence of Cupressaceae sensu lato since the Triassic in Gondwana. Key words: Barreal-Calingasta, Sorocayense Group, Voltziaceae, Cupressaceae sensu lato. RESUMO – Neste trabalho são descritos lenhos, folhas e ramos para a subclasse Pinidae (Voltziales e Pinales) da Formação Cortaderita (Triássico Médio). Os materiais foram encontrados nas quebradas La Cortaderita e La Tinta, na área de Barreal, bacia de Barreal-Calingasta, precordillera de San Juan, Argentina. Foram reconhecidas folhas correspondentes a três espécies do gênero Heidiphyllum (Voltziales), ramos folhosos atribuídos a Elatocladus planus (Pinales) e lenhos permineralizados designados como Cupressinoxylon zamunerae sp. nov. (Pinales). As duas últimas espécies têm características que permitem atribuí-los para a família Cupressaceae sensu lato. Por isso, neste estudo mostra-se que correspondem a mesma entidade biológica. Assim é descrito com o nome da nova espécie de Cupressinoxylon. Esta reconstrução é baseada em critérios de reforço mútuo e de ocorrência afi ns, além de características morfológicas e anatômicas. Desta forma, Cupressinoxylon zamunerae sp. nov. é interpretada como uma árvore de aproximadamente 13 m de altura, muito ramifi cada, com ramos da penúltima ordem com folhas escamiformes, e ramos de última ordem alternados, com folhas sésseis, uninerveadas, lineares a aciculares distribuídas helicoidalmente. Esses registros confi rmam a presença de Cupressaceae sensu lato, desde o Triássico no Gondwana desde o Triássico. Palavras-chave: Barreal-Calingasta, Grupo Sorocayense, Voltziaceae, Cupressaceae sensu lato. 141 142 REVISTA BRASILEIRA DE PALEONTOLOGIA, 18(1), 2015 INTRODUCCIÓN los leños registrados en la Argentina. En general, se acepta que las Voltziales y las Pinales compartían el mismo hábito Las comunidades arbóreas triásicas del Sudoeste de arbóreo, profusamente ramificado (Taylor et al., 2009). Gondwana estaban dominadas por Corystospermales Anatómicamente, el xilema secundario de las Voltziales (“Pteridospermas” o helechos con semilla) dejando a las triásicas y de las Pinales podría presentar similitudes. De coníferas un rol muy subordinado, que recién sería más hecho, en otras regiones del Gondwana, se ha sugerido importante a partir del Triásico Superior y el Jurásico una reconstrucción en la que se vincula a Heidiphyllum/ Inferior y Medio, precisamente gracias a la declinación de Telemachus con troncos permineralizados, de características las corystospermas (Artabe et al., 2007). semejantes a los leños de las podocarpáceas (Axsmith et al., El registro triásico de las coníferas en Sudamérica incluye 1998; Escapa et al., 2010; Bomfl eur et al., 2013). leños permineralizados, impresiones-compresiones de hojas, La necesidad de avanzar en el conocimiento de las ramas foliosas y conos, y granos de polen; asignados a los reconstrucciones de las coníferas triásicas, radica en órdenes Voltziales y Pinales (familias Cheirolepidiaceae, que es en dicho período donde se produce el origen y la Podocarpaceae, Araucariaceae, Cupressaceae sensu lato) diversifi cación de los clados modernos, puesto que para (Zamuner et al., 2001; Artabe et al., 2003). el Jurásico ya encontramos representantes de cada familia Actualmente se conocen para el Sistema Triásico (excepto por las Pinaceae) bien diferenciados (Stewart argentino las siguientes especies de leños asignadas a las & Rothwell, 1993). A la hora de vincular los diferentes coníferas: Araucariaceae, Agathoxylon protoaraucana órganos de coníferas triásicas en una reconstrucción, existen (Brea) Gnaedinger & Herbst, 2009, Agathoxylon varias difi cultades, e.g. el registro de los conos micro y lamaibandianus Crisafulli & Herbst, 2011, Agathoxylon megasporangiados es escaso o de difícil interpretación y dallonii (Boureau) Crisafulli & Herbst, 2010, Agathoxylon los leños de corystospermas en estado disperso son muy amraparense (Sah & Jain) Crisafulli & Herbst, 2011; semejantes anatómicamente a los de las coníferas. La alta Podocarpaceae, Podocarpoxylon tikiense Ram-Awatar & diversidad de leños permineralizados asignados a Pinales en Rajanikanth, 2007, Podocarpoxylon indicum (Bhardwaj) el Sistema Triásico de la Argentina, en comparación con la Bose & Maheshwari, 1974, Podocarpoxylon paralatifolium ausencia de conos micro y/o megasporangiados de Pinales, Vozenin-Serra & Grant-Mackie, 1996; Cupressaceae s.l., nos permite deducir que: (i) algunos ejemplares podrían Protojuniperoxylon ischigualastense Bonetti emend. Bodnar corresponder al grupo dominante de los bosques triásicos, & Artabe, 2007; y cuatro especies de familia incierta: i.e. Corystospermales; (ii) ciertas formas deberían pertenecer Protocircoporoxylon marianaensis Zamuner & Artabe, a la voltzial Heidiphyllum/Telemachus; (iii) diferencias en la 1994, Agathoxylon (= Araucarioxylon) sp. A. (Zamuner, preservación han permitido la fosilización de leños de Pinales 1992), Protophyllocladoxylon sp. (Crisafulli & Herbst, pero no así sus estructuras reproductivas. 2011), Prototaxoxylon intertrappeum Prakash & Srivastava, Para resolver algunas de estos interrogantes, los registros 1961 y Taxaceoxylon sp. (Caminos et al., 1995). Aunque las fl orísticos triásicos de la Formación Cortaderita (Groeber & dos últimas fueron asignadas a las Taxaceae (Zamuner et Stipanicic, 1953), Grupo Sorocayense, Cuenca de Barreal- al., 2001; Crisafulli & Herbst, 2010), los caracteres en los Calingasta, San Juan, centro-oeste de la Argentina (Figura 1) que se basa esa afi rmación probablemente sean productos de presentan una gran ventaja, por la continuidad vertical y la degradación de la pared de las células xilemáticas o que horizontal de los estratos plantíferos, la gran diversidad correspondan a leño de compresión. taxonómica y de condiciones tafonómicas, y la presencia de Por otra parte, también son abundantes los registros una serie de bosques fosilizados in situ correspondiendo a de formas foliares de coníferas: Heidiphyllum elongatum tafocenosis autóctonas (Artabe et al., 2001; Bodnar, 2010). (Morris) Retallack, 1981, entre las Voltziales; y Elatocladus Para la Formación Cortaderita, la primera cita de un planus (Feistmantel) Seward, 1919, E. australis Frenguelli, leño fósil de conífera corresponde a Menéndez (1956) quien 1944, Rissikia media (Tenison Woods) Townrow, 1967 y erigió la especie Protophyllocladoxylon cortaderitaense Pagiophyllum sp., entre las Pinales (Artabe et al., 2007). sobre la base de material encontrado en la sección superior En cuanto a las estructuras reproductivas, son frecuentes de la unidad, en la localidad de Quebrada de la Tinta. los conos megasporangiados de las Voltziales: Telemachus Posteriormente, Lutz & Herbst (1992) describieron troncos lignosus Retallack, 1981 y T. elongatus Anderson, 1978, de una corystosperma, hallados en la sección inferior de la Dordrechtites sp. (Spalletti et al., 1991; Arce & Lutz, 2010; Formación Cortaderita, en la Quebrada homónima, y los Morel
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